treatments-xml/data/03/D7/98/03D79875FFFEFFB2FF4EFDCC482804CD.xml

<document id="56BE297329906EBD245F41BC8CB309DE" ID-DOI="10.5281/zenodo.281817" ID-GBIF-Dataset="9c482ced-fa54-4bb6-a31c-2e8f0b205f40" ID-ISSN="1175-5326" ID-Zenodo-Dep="281817" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1460501071456" checkinUser="plazi" docAuthor="Hamidy, Amir, Matsui, Masafumi, Nishikawa, Kanto &amp; Belabut, Daicus M." docDate="2012" docId="03D79875FFFEFFB2FF4EFDCC482804CD" docLanguage="en" docName="zt03398p039.pdf" docOrigin="Zootaxa 3398" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Leptobrachium ingeri Hamidy, Matsui, Nishikawa &amp; Belabut, 2012, sp. nov." docType="treatment" docVersion="9" lastPageNumber="34" masterDocId="FFEEE00DFFF8FFBEFFD9FFA64B0E033F" masterDocTitle="Detection of cryptic taxa in Leptobrachium nigrops (Amphibia, Anura, Megophryidae), with description of two new species" masterLastPageNumber="39" masterPageNumber="22" pageNumber="28" updateTime="1698311801674" updateUser="plazi">
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<mods:title id="16C2A19A2924A94F1F748D04F4BF9EA3">Detection of cryptic taxa in Leptobrachium nigrops (Amphibia, Anura, Megophryidae), with description of two new species</mods:title>
</mods:titleInfo>
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<mods:namePart id="0626A2A8C27959C3E30B99BA33790D0D">Hamidy, Amir</mods:namePart>
</mods:name>
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<mods:namePart id="72256B328E02BC720E64FCE7F8EB8994">Matsui, Masafumi</mods:namePart>
</mods:name>
<mods:name id="ADEC2D3828A4872EF412D6A25637EA7B" type="personal">
<mods:role id="90D744C437CD59E31AB00D9877C9B4FF">
<mods:roleTerm id="05126BEF38438B1E169A898B1B99A922">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="76E15942356452C9C25BFC77A4D84715">Nishikawa, Kanto</mods:namePart>
</mods:name>
<mods:name id="CBF67AF4AE42BD88E4D15D662BDC213A" type="personal">
<mods:role id="D487914F89DA98E29CC8B5A515D7F692">
<mods:roleTerm id="C2D661C8C9A0A92A0E4341186C31D2B9">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="6EFD9CA9A4CCF4305AEE86959A8B4E99">Belabut, Daicus M.</mods:namePart>
</mods:name>
<mods:typeOfResource id="4E11B749DA036E3CE46DF97D51B47367">text</mods:typeOfResource>
<mods:relatedItem id="0D00ED40505CE0D9348EEA000D1BAF9A" type="host">
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<mods:title id="F6BBE512549EE76D49F46ED83B268CD5">Zootaxa</mods:title>
</mods:titleInfo>
<mods:part id="714745FAD430057E64A77B382026123D">
<mods:date id="860C6FC6157FD092475C833B7C8B5785">2012</mods:date>
<mods:detail id="046DF32EBA153D76E874C1A90EDFF98B" type="volume">
<mods:number id="84138114B01C550AD6D92C76CCE7C8AF">3398</mods:number>
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<mods:classification id="13CE6E42191292B143569979CF7602BD">journal article</mods:classification>
<mods:identifier id="57A55E8028347E6AED6C63F4FEDD18AD" type="DOI">10.5281/zenodo.281817</mods:identifier>
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<mods:identifier id="D4EFE99520E2BCD960A7BF2B243AFB90" type="ISSN">1175-5326</mods:identifier>
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<treatment id="03D79875FFFEFFB2FF4EFDCC482804CD" ID-DOI="http://doi.org/10.5281/zenodo.6180104" ID-GBIF-Taxon="119521421" ID-Zenodo-Dep="6180104" LSID="urn:lsid:zoobank.org:act:1239011D-682D-486D-B095-FB8181076EA9" httpUri="http://treatment.plazi.org/id/03D79875FFFEFFB2FF4EFDCC482804CD" lastPageId="12" lastPageNumber="34" pageId="6" pageNumber="28">
<subSubSection id="C3647AE8FFFEFFB8FF4EFDCC497101F7" pageId="6" pageNumber="28" type="nomenclature">
<paragraph id="8BC12963FFFEFFB8FF4EFDCC490A01BB" blockId="6.[151,516,618,644]" box="[151,516,618,644]" pageId="6" pageNumber="28">
<heading id="D0899E0FFFFEFFB8FF4EFDCC490A01BB" bold="true" box="[151,516,618,644]" fontSize="11" level="1" pageId="6" pageNumber="28" reason="1">
<emphasis id="B90AF571FFFEFFB8FF4EFDCC490A01BB" bold="true" box="[151,516,618,644]" pageId="6" pageNumber="28">
<taxonomicName id="4C7E52E0FFFEFFB8FF4EFDCC4AAE01BB" ID-CoL="3TKC5" box="[151,416,618,644]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="6" pageNumber="28" phylum="Chordata" rank="species" species="ingeri" status="sp. nov.">
<emphasis id="B90AF571FFFEFFB8FF4EFDCC4AAE01BB" bold="true" box="[151,416,618,644]" italics="true" pageId="6" pageNumber="28">Leptobrachium ingeri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFFEFFB8FE7EFDCC490A01BB" box="[423,516,618,644]" pageId="6" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BC12963FFFEFFB8FF4EFD17497101F7" blockId="6.[151,639,689,712]" box="[151,639,689,712]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF4EFD174A1C01F7" bold="true" box="[151,274,689,712]" pageId="6" pageNumber="28">Synonymy:</emphasis>
<taxonomicName id="4C7E52E0FFFEFFB8FECEFD144AF201F8" authority="Inger 1966" authorityName="Inger" authorityYear="1966" box="[279,508,689,711]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="6" pageNumber="28" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFFEFFB8FECEFD144A7001F8" box="[279,382,690,711]" italics="true" pageId="6" pageNumber="28">L. nigrops</emphasis>
: 
<bibRefCitation id="EFEF5492FFFEFFB8FE55FD144AF201F8" author="Inger" box="[396,508,689,711]" pageId="6" pageNumber="28" refString="Inger, R. F. (1966) The systematics and zoogeography of the Amphibia of Borneo. Fieldiana: Zoology, 52, 1 - 402." type="journal article" year="1966">Inger 1966</bibRefCitation>
</taxonomicName>
, p. 37 (part).
</paragraph>
</subSubSection>
<subSubSection id="C3647AE8FFFEFFB8FF4EFD524A5E074B" pageId="6" pageNumber="28" type="materials_examined">
<paragraph id="8BC12963FFFEFFB8FF4EFD524A20006B" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF4EFD524A050033" bold="true" box="[151,267,756,780]" pageId="6" pageNumber="28">
<typeStatus id="54C597C1FFFEFFB8FF4EFD524A080033" box="[151,262,756,780]" pageId="6" pageNumber="28" type="holotype">Holotype</typeStatus>
.
</emphasis>
<collectionCode id="ED6FB1A6FFFEFFB8FECAFD534A6F0033" box="[275,353,757,780]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
53848, an adult female collected from Matang Wildlife Centre (
<geoCoordinate id="EE4A4FA4FFFEFFB8FBE4FD524FC30033" box="[1085,1229,756,780]" direction="north" orientation="latitude" pageId="6" pageNumber="28" precision="15" value="1.6094445">01°36’34’’N</geoCoordinate>
, 
<geoCoordinate id="EE4A4FA4FFFEFFB8FB03FD524E7A0033" box="[1242,1396,756,780]" direction="east" orientation="longitude" pageId="6" pageNumber="28" precision="15" value="110.16">110°09’36’’E</geoCoordinate>
, 
<quantity id="4C868486FFFEFFB8FA59FD524BA30010" metricMagnitude="1" metricUnit="m" metricValue="6.0" pageId="6" pageNumber="28" unit="m" value="60.0">60 m</quantity>
a.s.l.), Kuching Division, 
<collectingCountry id="F36969F3FFFEFFB8FE05FCB149310010" box="[476,575,791,815]" name="Malaysia" pageId="6" pageNumber="28">Sarawak</collectingCountry>
, Borneo, 
<collectingCountry id="F36969F3FFFEFFB8FD77FCB148190010" box="[686,791,791,815]" name="Malaysia" pageId="6" pageNumber="28">Malaysia</collectingCountry>
, collected by K. Eto and K. Nishikawa at 20:00 h on 
<date id="FFC00FA3FFFEFFB8FA59FCB14A25006B" pageId="6" pageNumber="28" value="2010-08-30">30 August 2010</date>
.
</paragraph>
<paragraph id="8BC12963FFFEFFB8FF10FCC64A5F0738" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF10FCC64A460047" bold="true" box="[201,328,864,888]" pageId="6" pageNumber="28">
<typeStatus id="54C597C1FFFEFFB8FF10FCC64A4D0047" box="[201,323,864,888]" pageId="6" pageNumber="28" type="paratype">Paratypes</typeStatus>
.
</emphasis>
SRC 0 0 0 1 (former 
<collectionCode id="ED6FB1A6FFFEFFB8FDFFFCC7497A0047" box="[550,628,865,888]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
53833) and 
<collectionCode id="ED6FB1A6FFFEFFB8FCD8FCC748410047" box="[769,847,865,888]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
53834, two adult males, and 
<collectionCode id="ED6FB1A6FFFEFFB8FB40FCC74FE90047" box="[1177,1255,865,888]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
53832, an adult female, all collected from Santubong (
<geoCoordinate id="EE4A4FA4FFFEFFB8FD94FC2249D300A3" box="[589,733,900,924]" direction="north" orientation="latitude" pageId="6" pageNumber="28" precision="15" value="1.7436111">01°44’37’’N</geoCoordinate>
, 
<geoCoordinate id="EE4A4FA4FFFEFFB8FD33FC22488C00A3" box="[746,898,900,924]" direction="east" orientation="longitude" pageId="6" pageNumber="28" precision="15" value="110.31972">110°19’11’’E</geoCoordinate>
, 
<quantity id="4C868486FFFEFFB8FC49FC2248C500A3" box="[912,971,900,924]" metricMagnitude="1" metricUnit="m" metricValue="6.0" pageId="6" pageNumber="28" unit="m" value="60.0">60 m</quantity>
a.s.l.), Kuching Division, 
<collectingCountry id="F36969F3FFFEFFB8FB20FC224E5200A3" box="[1273,1372,900,924]" name="Malaysia" pageId="6" pageNumber="28">Sarawak</collectingCountry>
, Borneo, 
<collectingCountry id="F36969F3FFFEFFB8FF16FC014A380080" box="[207,310,935,959]" name="Malaysia" pageId="6" pageNumber="28">Malaysia</collectingCountry>
on 
<date id="FFC00FA3FFFEFFB8FEBDFC0149100080" box="[356,542,935,960]" pageId="6" pageNumber="28" value="2010-08-29">29 August 2010</date>
, collectors same as for 
<typeStatus id="54C597C1FFFEFFB8FCF7FC01489A0080" box="[814,916,935,959]" pageId="6" pageNumber="28" type="holotype">holotype</typeStatus>
; 
<collectionCode id="ED6FB1A6FFFEFFB8FC78FC0F48D000FF" box="[929,990,937,960]" httpUri="http://grbio.org/cool/b9gz-izvg" name="Museum Zoologicum Bogoriense" pageId="6" pageNumber="28">MZB</collectionCode>
Amp 11790, an adult female collected from marshy area of Tanjung Padang, Belitung, 
<collectingCountry id="F36969F3FFFEFFB8FD61FC6A482600DB" box="[696,808,972,996]" name="Indonesia" pageId="6" pageNumber="28">Indonesia</collectingCountry>
, collected by D. Gower, A. Hamidy, and K. Sanders on 
<date id="FFC00FA3FFFEFFB8FF4EFC494A420738" box="[151,332,1007,1031]" pageId="6" pageNumber="28" value="2005-12">December 2005</date>
.
</paragraph>
<paragraph id="8BC12963FFFEFFB8FF10FBB54A5E074B" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF10FBB54AB30713" bold="true" box="[201,445,1043,1068]" pageId="6" pageNumber="28">Referred specimens.</emphasis>
<collectionCode id="ED6FB1A6FFFEFFB8FE1FFBB3491A0713" box="[454,532,1045,1068]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
12087 and 
<collectionCode id="ED6FB1A6FFFEFFB8FD47FBB349E20713" box="[670,748,1045,1068]" httpUri="http://grbio.org/cool/jxgt-7i97" name="Kyoto University, Graduate School of Human and Environmental Studies" pageId="6" pageNumber="28">KUHE</collectionCode>
12099, two adult males, collected from Niah National Park (
<geoCoordinate id="EE4A4FA4FFFEFFB8FF46FB9E4A0D076F" box="[159,259,1080,1104]" direction="north" orientation="latitude" pageId="6" pageNumber="28" precision="925" value="3.8166666">03°49’N</geoCoordinate>
, 
<geoCoordinate id="EE4A4FA4FFFEFFB8FECBFB9E4A71076F" box="[274,383,1080,1104]" direction="east" orientation="longitude" pageId="6" pageNumber="28" precision="925" value="113.75">113°45’E</geoCoordinate>
, ca. 
<quantity id="4C868486FFFEFFB8FE6EFB9E4AFD076F" box="[439,499,1080,1104]" metricMagnitude="1" metricUnit="m" metricValue="5.0" pageId="6" pageNumber="28" unit="m" value="50.0">50 m</quantity>
a.s.l.), Miri Division, 
<collectingCountry id="F36969F3FFFEFFB8FD21FB9E4855076F" box="[760,859,1080,1104]" name="Malaysia" pageId="6" pageNumber="28">Sarawak</collectingCountry>
, Borneo, 
<collectingCountry id="F36969F3FFFEFFB8FC15FB9E4F38076F" box="[972,1078,1080,1104]" name="Malaysia" pageId="6" pageNumber="28">Malaysia</collectingCountry>
, collected by M. Matsui on 
<date id="FFC00FA3FFFEFFB8FA59FB9E4A42074B" pageId="6" pageNumber="28" value="1990-12-26">26 December 1990</date>
.
</paragraph>
</subSubSection>
<subSubSection id="C3647AE8FFFEFFB8FF10FB264E350783" pageId="6" pageNumber="28" type="etymology">
<paragraph id="8BC12963FFFEFFB8FF10FB264E350783" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF10FB264A4107A7" bold="true" box="[201,335,1152,1176]" pageId="6" pageNumber="28">Etymology.</emphasis>
The specific name is dedicated to Dr. Robert F. Inger of Field Museum, Chicago, who was the first to notice morphological differences between populations of 
<taxonomicName id="4C7E52E0FFFEFFB8FCE1FB0348A40783" box="[824,938,1189,1212]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="6" pageNumber="28" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFFEFFB8FCE1FB0348A40783" box="[824,938,1189,1212]" italics="true" pageId="6" pageNumber="28">L. nigrops</emphasis>
</taxonomicName>
from Borneo and Malay Peninsula.
</paragraph>
</subSubSection>
<subSubSection id="C3647AE8FFFEFFB8FF10FB614F4D06AB" pageId="6" pageNumber="28" type="diagnosis">
<paragraph id="8BC12963FFFEFFB8FF10FB614F4D06AB" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF10FB614A4D07DF" bold="true" box="[201,323,1223,1248]" pageId="6" pageNumber="28">Diagnosis.</emphasis>
A small-sized 
<taxonomicName id="4C7E52E0FFFEFFB8FE37FB6E499707DF" box="[494,665,1224,1248]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="6" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="B90AF571FFFEFFB8FE37FB6E499707DF" box="[494,665,1224,1248]" italics="true" pageId="6" pageNumber="28">Leptobrachium</emphasis>
</taxonomicName>
of the 
<taxonomicName id="4C7E52E0FFFEFFB8FD32FB6F485307DF" box="[747,861,1225,1248]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="6" pageNumber="28" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFFEFFB8FD32FB6F485307DF" box="[747,861,1225,1248]" italics="true" pageId="6" pageNumber="28">L. nigrops</emphasis>
</taxonomicName>
group, with SVL of 
<quantity id="4C868486FFFEFFB8FB92FB6E4FE107DF" box="[1099,1263,1224,1248]" metricMagnitude="-2" metricUnit="m" metricValue="4.32" metricValueMax="4.71" metricValueMin="3.9299999999999997" pageId="6" pageNumber="28" unit="mm" value="43.2" valueMax="47.1" valueMin="39.3">39.3–47.1 mm</quantity>
in females and 
<quantity id="4C868486FFFEFFB8FF4EFB4A4A32063B" box="[151,316,1260,1284]" metricMagnitude="-2" metricUnit="m" metricValue="3.2700000000000005" metricValueMax="3.6799999999999997" metricValueMin="2.8600000000000003" pageId="6" pageNumber="28" unit="mm" value="32.7" valueMax="36.8" valueMin="28.6">28.6–36.8 mm</quantity>
in males; finger tip sharply pointed; toe web relatively well developed; inner metatarsal tubercle large; dorsum brown with distinct dark brown markings; tympanum usually at least upper one-third covered by dark brown marking; body laterally with black spots scattered from axilla to groin; belly and limbs heavily blotched ventrally; limbs, including fingers and toes, distinctly barred dorsally; large dark blotches at groin continuing to ventral and posterior sides of thigh; femoral gland in a large white blotch.
</paragraph>
</subSubSection>
<subSubSection id="C3647AE8FFFEFFB2FF10FA39482804CD" lastPageId="12" lastPageNumber="34" pageId="6" pageNumber="28" type="description">
<paragraph id="8BC12963FFFEFFB8FF10FA39493705C3" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">
<emphasis id="B90AF571FFFEFFB8FF10FA3948060687" bold="true" box="[201,776,1439,1464]" pageId="6" pageNumber="28">
Description of 
<typeStatus id="54C597C1FFFEFFB8FEA5FA064AED0687" box="[380,483,1440,1464]" pageId="6" pageNumber="28" type="holotype">holotype</typeStatus>
(measurements in mm).
</emphasis>
Small sized (SVL 44.6); habitus moderately stocky, body tapering to groin, head broad and depressed, slightly longer (HL 19.6: 43.9% SVL) than wide (HW 18.0: 40.4% SVL); snout obtusely pointed, rounded triangle from above, truncate in profile, projecting beyond lower jaw; eye large, obviously projecting from sides of head, slightly smaller (EL 6.8: 15.3% SVL) than snout (SL 7.3: 16.4% SVL); canthus rostralis sharp, loreal region oblique, moderately concave; nostril lateral, below canthus, closer to snout (S-NL 3.9: 8.7% SVL) than to eye (N-EL 4.1: 9.2% SVL); internarial distance (
<collectingCountry id="F36969F3FFFEFFB8FBBEF9F24F950553" box="[1127,1179,1620,1644]" name="India" pageId="6" pageNumber="28">IND</collectingCountry>
3.4: 7.6% SVL) much narrower than interorbital distance, (IOD 5.7: 12.8% SVL), latter slightly narrower than upper eyelid (UEW 6.0: 13.5% SVL); pineal spot absent; tympanum visible, diameter (TD 3.2: 7.2% SVL) almost half that of eye and separated from eye by about two-third of its diameter (T-EL 2.2: 4.9% SVL); no vomerine teeth; tongue heart-shaped, without papillae, notched posteriorly.
</paragraph>
<paragraph id="8BC12963FFFEFFB8FF10F8AE49DE048F" blockId="6.[151,1437,756,1968]" pageId="6" pageNumber="28">Forelimb slender and long (FLL 31.3: 63.7% SVL), about three-fifth of hindlimb; fingers moderately slender, unwebbed; first finger (1FL 5.9: 10.7% SVL) slightly longer than fourth and second, third much longer (3FL 8.9: 16.2% SVL); finger tip sharply pointed, slightly hooked, not swollen; inner palmar tubercle large (IPTL 2.5: 4.5% SVL), not extending onto first metacarpal, and outer palmar tubercle smaller (OPTL 2.0: 3.6% SVL); subarticular tubercles indistinct, replaced by low callous tissue.</paragraph>
<caption id="DF0179EBFFFFFFB9FF4EF83D49A504D0" httpUri="https://zenodo.org/record/281820/files/figure.png" pageId="7" pageNumber="29" targetBox="[332,1255,176,1926]" targetPageId="7">
<paragraph id="8BC12963FFFFFFB9FF4EF83D49A504D0" blockId="7.[151,1436,1947,2031]" pageId="7" pageNumber="29">
<emphasis id="B90AF571FFFFFFB9FF4EF83D4A18048E" bold="true" box="[151,278,1947,1970]" pageId="7" pageNumber="29">FIGURE 3.</emphasis>
Three species of the 
<taxonomicName id="4C7E52E0FFFFFFB9FE24F83A49E0048D" box="[509,750,1948,1970]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="7" pageNumber="29" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFFFFFB9FE24F83A49E0048D" box="[509,750,1948,1970]" italics="true" pageId="7" pageNumber="29">Leptobrachium nigrops</emphasis>
</taxonomicName>
group in life. (A) A female of 
<taxonomicName id="4C7E52E0FFFFFFB9FB98F83B4FA5048D" box="[1089,1195,1949,1970]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="7" pageNumber="29" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFFFFFB9FB98F83B4FA5048D" box="[1089,1195,1949,1970]" italics="true" pageId="7" pageNumber="29">L. nigrops</emphasis>
</taxonomicName>
from Malay Peninsula (KUHE 42243); (B) A female paratype of 
<taxonomicName id="4C7E52E0FFFFFFB9FD8EF81D49BE04EF" box="[599,688,1979,2000]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="7" pageNumber="29" phylum="Chordata" rank="species" species="ingeri" status="sp. nov.">
<emphasis id="B90AF571FFFFFFB9FD8EF81D49BE04EF" box="[599,688,1979,2000]" italics="true" pageId="7" pageNumber="29">L. ingeri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFFFFFB9FD63F81D480304EF" box="[698,781,1979,2000]" pageId="7" pageNumber="29" rank="species">
<emphasis id="B90AF571FFFFFFB9FD63F81D480304EF" bold="true" box="[698,781,1979,2000]" pageId="7" pageNumber="29">sp. nov.</emphasis>
</taxonomicNameLabel>
from Santubong (KUHE 53832); (C) A female holotype of 
<taxonomicName id="4C7E52E0FFFFFFB9FA5EF81D4A1A04D0" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="7" pageNumber="29" phylum="Chordata" rank="species" species="kanowitense" status="sp. nov.">
<emphasis id="B90AF571FFFFFFB9FA5EF81D4A1A04D0" italics="true" pageId="7" pageNumber="29">L. kanowitense</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFFFFFB9FEC3F87C4A6504D0" box="[282,363,2010,2031]" pageId="7" pageNumber="29" rank="species">
<emphasis id="B90AF571FFFFFFB9FEC3F87C4A6504D0" bold="true" box="[282,363,2010,2031]" pageId="7" pageNumber="29">sp. nov.</emphasis>
</taxonomicNameLabel>
from Kanowit (KUHE 42575).
</paragraph>
</caption>
<caption id="DF0179EBFFF0FFB6FF4EFB6A4E430620" httpUri="https://zenodo.org/record/281821/files/figure.png" pageId="8" pageNumber="30" targetBox="[160,1415,221,1181]" targetPageId="8">
<paragraph id="8BC12963FFF0FFB6FF4EFB6A4E430620" blockId="8.[151,1436,1228,1311]" pageId="8" pageNumber="30">
<emphasis id="B90AF571FFF0FFB6FF4EFB6A4BF407DC" bold="true" box="[151,250,1228,1251]" pageId="8" pageNumber="30">FIGURE</emphasis>
4. Ventral views in males (top row) and females (bottom row) of (A) 
<taxonomicName id="4C7E52E0FFF0FFB6FC0BFB6B4F3407DD" box="[978,1082,1229,1250]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="8" pageNumber="30" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF0FFB6FC0BFB6B4F3407DD" box="[978,1082,1229,1250]" italics="true" pageId="8" pageNumber="30">L. nigrops</emphasis>
</taxonomicName>
: KUHE 15658 (male) and KUHE 42243 (female); (B) 
<taxonomicName id="4C7E52E0FFF0FFB6FEABFB4A4AC3063E" box="[370,461,1260,1281]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="8" pageNumber="30" phylum="Chordata" rank="species" species="ingeri" status="sp. nov.">
<emphasis id="B90AF571FFF0FFB6FEABFB4A4AC3063E" box="[370,461,1260,1281]" italics="true" pageId="8" pageNumber="30">L. ingeri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFF0FFB6FE0FFB4D4924063F" box="[470,554,1259,1280]" pageId="8" pageNumber="30" rank="species">
<emphasis id="B90AF571FFF0FFB6FE0FFB4D4924063F" bold="true" box="[470,554,1259,1280]" pageId="8" pageNumber="30">sp. nov.</emphasis>
</taxonomicNameLabel>
: a paratype (KUHE 53834: male) and the holotype (KUHE 53848: female); (C) 
<taxonomicName id="4C7E52E0FFF0FFB6FA5EFB4A4A1A0620" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="8" pageNumber="30" phylum="Chordata" rank="species" species="kanowitense" status="sp. nov.">
<emphasis id="B90AF571FFF0FFB6FA5EFB4A4A1A0620" italics="true" pageId="8" pageNumber="30">L. kanowitense</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFF0FFB6FEC3FAAC4A650620" box="[282,363,1290,1311]" pageId="8" pageNumber="30" rank="species">
<emphasis id="B90AF571FFF0FFB6FEC3FAAC4A650620" bold="true" box="[282,363,1290,1311]" pageId="8" pageNumber="30">sp. nov.</emphasis>
</taxonomicNameLabel>
: a paratype (KUHE42574: male) and the holotype (KUHE 42575: female). Scale indicates 5 mm.
</paragraph>
</caption>
<paragraph id="8BC12963FFF0FFB6FF10FAE14FE70527" blockId="8.[151,1436,1351,2028]" pageId="8" pageNumber="30">
Hindlimb slender and relatively short (HLL 65.6: 119.2% SVL); heels not meeting when legs held at right angles to body; tibia distinctly longer (TL 20.0: 36.3% SVL) than foot (FL 17.0: 30.9% SVL); tibiotarsal articulation of adpressed limb reaching to middle of tympanum; third toe longer than fifth; toe tips similar to those of fingers; toe webs rather well developed, webbing formula 
<date id="FFC00FA3FFF0FFB6FC86FA11489506F1" box="[863,923,1462,1486]" pageId="8" pageNumber="30">I 1–1</date>
1/ 
<date id="FFC00FA3FFF0FFB6FC72FA6348C106F1" box="[939,975,1463,1491]" pageId="8" pageNumber="30">2 II</date>
1/4–2 3/ 
<date id="FFC00FA3FFF0FFB6FBC5FA634F4706F1" box="[1052,1097,1463,1491]" pageId="8" pageNumber="30">4 III</date>
1 1/4–3 2/ 
<date id="FFC00FA3FFF0FFB6FB7AFA634FDC06F1" box="[1187,1234,1463,1491]" pageId="8" pageNumber="30">3 IV</date>
3 2/3–1 1/ 
<date id="FFC00FA3FFF0FFB6FAF5FA634E5C06F1" box="[1324,1362,1463,1491]" pageId="8" pageNumber="30">2 V</date>
; inner metatarsal tubercle oval, low but large (IMTL 2.2: 4.9% SVL) more than half of first toe (1TOEL 3.2:7.2%); outer metatarsal tubercle absent; subarticular tubercles obscure, elongate, replaced by low callous tissue.
</paragraph>
<paragraph id="8BC12963FFF0FFB6FF10F9824BF605F3" blockId="8.[151,1436,1351,2028]" pageId="8" pageNumber="30">Skin above nearly smooth, scattered with minute granules on top of head, especially around interorbital region, granules denser on upper eyelid; ventrum weakly granular, particularly on belly; low supratympanic ridge from posterior corner of eye to axilla; low dermal ridges in longitudinal rows on upper arm; flat pectoral gland at medial border of axilla behind arm insertion; femoral gland small, concentrated in one cluster on right side, but diffused on left side.</paragraph>
<paragraph id="8BC12963FFF0FFB6FF10F97E49D204D3" blockId="8.[151,1436,1351,2028]" pageId="8" pageNumber="30">
<emphasis id="B90AF571FFF0FFB6FF10F97E4A2C05CE" bold="true" box="[201,290,1752,1777]" pageId="8" pageNumber="30">Colour.</emphasis>
In life, dorsally brown with dark brown blotches in interorbital region, medial one followed by continuous longitudinal band and lateral ones by more or less interrupted bands; dorsal brown laterally with distinct dark spots fading to lighter towards belly; loreal region, anterior to eye, covered by dark brown; upper onethird of tympanum and supratympanic ridge covered by dark band; groin marked with irregular dark blotches; ventrum cream heavily blotched all-over; limbs with distinct black bars dorsally and irregular dark blotches ventrally; posterior thigh banded by black bars, banded pattern continuing to posterior flank; iris totally black, surrounded by light blue sclera visible in maximally opened eye. In preservative, aspects of the colour pattern remain, but dorsal brown colour has been darkened.
</paragraph>
<caption id="DF0179EBFFF2FFB4FF4EF8124E2E04D6" httpUri="https://zenodo.org/record/281822/files/figure.png" pageId="10" pageNumber="32" targetBox="[186,1408,193,1957]" targetPageId="10">
<paragraph id="8BC12963FFF2FFB4FF4EF8124E2E04D6" blockId="10.[151,1436,1972,2026]" pageId="10" pageNumber="32">
<emphasis id="B90AF571FFF2FFB4FF4EF8124A1D04F5" bold="true" box="[151,275,1972,1995]" pageId="10" pageNumber="32">FIGURE 5.</emphasis>
Ventral sides of thigh (top row), hand (middle row), and foot (bottom row) of (A) 
<taxonomicName id="4C7E52E0FFF2FFB4FBB8F8104FC604F4" box="[1121,1224,1974,1995]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="10" pageNumber="32" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF2FFB4FBB8F8104FC604F4" box="[1121,1224,1974,1995]" italics="true" pageId="10" pageNumber="32">L. nigrops</emphasis>
</taxonomicName>
(KUHE 42243), (B) the holotype of 
<taxonomicName id="4C7E52E0FFF2FFB4FEEFF8724A8304D6" box="[310,397,2004,2025]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="10" pageNumber="32" phylum="Chordata" rank="species" species="ingeri" status="sp. nov.">
<emphasis id="B90AF571FFF2FFB4FEEFF8724A8304D6" box="[310,397,2004,2025]" italics="true" pageId="10" pageNumber="32">L. ingeri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFF2FFB4FE4DF8724AEA04D6" box="[404,484,2004,2025]" pageId="10" pageNumber="32" rank="species">
<emphasis id="B90AF571FFF2FFB4FE4DF8724AEA04D6" bold="true" box="[404,484,2004,2025]" pageId="10" pageNumber="32">sp. nov.</emphasis>
</taxonomicNameLabel>
(KUHE 53848), and (C) the holotype of 
<taxonomicName id="4C7E52E0FFF2FFB4FC5EF8724F1104D6" box="[903,1055,2003,2025]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="10" pageNumber="32" phylum="Chordata" rank="species" species="kanowitense" status="sp. nov.">
<emphasis id="B90AF571FFF2FFB4FC5EF8724F1104D6" box="[903,1055,2003,2025]" italics="true" pageId="10" pageNumber="32">L. kanowitense</emphasis>
</taxonomicName>
<emphasis id="B90AF571FFF2FFB4FBFCF8724F7804D6" bold="true" box="[1061,1142,2004,2025]" pageId="10" pageNumber="32">
<taxonomicNameLabel id="A239480AFFF2FFB4FBFCF8724F7804D6" box="[1061,1142,2004,2025]" pageId="10" pageNumber="32" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
(KUHE 42575).
</paragraph>
</caption>
<caption id="DF0179EBFFF3FFB5FF4EF95E4F500412" httpUri="https://zenodo.org/record/281823/files/figure.png" pageId="11" pageNumber="33" targetBox="[221,1371,238,1720]" targetPageId="11">
<paragraph id="8BC12963FFF3FFB5FF4EF95E4F500412" blockId="11.[151,1435,1784,1838]" pageId="11" pageNumber="33">
<emphasis id="B90AF571FFF3FFB5FF4EF95E4A1D0431" bold="true" box="[151,275,1784,1807]" pageId="11" pageNumber="33">FIGURE 6.</emphasis>
Spectrograms (above) and wave form (below) of advertisement call of (A) 
<taxonomicName id="4C7E52E0FFF3FFB5FBCCF95C4F730430" box="[1045,1149,1786,1807]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="11" pageNumber="33" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF3FFB5FBCCF95C4F730430" box="[1045,1149,1786,1807]" italics="true" pageId="11" pageNumber="33">L. nigrops</emphasis>
</taxonomicName>
(KUHE 15430) from Kuala Lumpur, Malay Peninsula and (B) 
<taxonomicName id="4C7E52E0FFF3FFB5FE2EF8BE49400412" box="[503,590,1816,1837]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="11" pageNumber="33" phylum="Chordata" rank="species" species="ingeri" status="sp. nov.">
<emphasis id="B90AF571FFF3FFB5FE2EF8BE49400412" box="[503,590,1816,1837]" italics="true" pageId="11" pageNumber="33">L. ingeri</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A239480AFFF3FFB5FD8DF8BE49AB0412" box="[596,677,1816,1837]" pageId="11" pageNumber="33" rank="species">
<emphasis id="B90AF571FFF3FFB5FD8DF8BE49AB0412" bold="true" box="[596,677,1816,1837]" pageId="11" pageNumber="33">sp. nov.</emphasis>
</taxonomicNameLabel>
(KUHE 53834) from Santubong, Sarawak.
</paragraph>
</caption>
<paragraph id="8BC12963FFF3FFB2FF10F8FF49230223" blockId="11.[151,1436,1881,2015]" lastBlockId="12.[151,1437,151,2034]" lastPageId="12" lastPageNumber="34" pageId="11" pageNumber="33">
<emphasis id="B90AF571FFF3FFB5FF10F8FF4A30044D" bold="true" box="[201,318,1881,1906]" pageId="11" pageNumber="33">Variation.</emphasis>
Morphological variations are shown in Table 3. The male 
<typeStatus id="54C597C1FFF3FFB5FC33F8FD4F44044C" box="[1002,1098,1883,1907]" pageId="11" pageNumber="33" type="paratype">paratype</typeStatus>
(KUHE 53834) is smaller in body size, and has an internal vocal sac and a pair of vocal sac openings. The female 
<typeStatus id="54C597C1FFF3FFB5FBB0F8D94FC704A8" box="[1129,1225,1919,1943]" pageId="11" pageNumber="33" type="paratype">paratype</typeStatus>
(KUHE 53832) is larger in body size with much darker throat than the 
<typeStatus id="54C597C1FFF3FFB5FD32F804485F0485" box="[747,849,1954,1978]" pageId="11" pageNumber="33" type="holotype">holotype</typeStatus>
. It has distinct white dots around vent and posterior thigh. The female 
<typeStatus id="54C597C1FFF3FFB5FEBDF8614ACA04E0" box="[356,452,1991,2015]" pageId="11" pageNumber="33" type="paratype">paratype</typeStatus>
from Belitung (MZB Amp 11790) is smaller than 
<typeStatus id="54C597C1FFF3FFB5FC24F8604F6D04E1" box="[1021,1123,1990,2014]" pageId="11" pageNumber="33" type="holotype">holotype</typeStatus>
. Although the color slightly faded, blotches on groin and posterior thigh can be still recognized. Specimens from Belitung have larger femoral gland and more rugose dorsal skin than specimens from Borneo. Like the 
<typeStatus id="54C597C1FFF4FFB2FC04FF1A4F4C03EB" box="[989,1090,188,212]" pageId="12" pageNumber="34" type="holotype">holotype</typeStatus>
, tibio-tarsal articulation in one female reaches only to middle of tympanum, but it reaches to points beyond anterior border of tympanum in the remaining one female and all males.
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF1EFE814BFA0103" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
<emphasis id="B90AF571FFF4FFB2FF1EFE814A04027F" bold="true" box="[199,266,295,320]" pageId="12" pageNumber="34">Calls.</emphasis>
The calls recorded at Santubong on 
<date id="FFC00FA3FFF4FFB2FD79FE81485F0200" box="[672,849,295,320]" pageId="12" pageNumber="34" value="2010-08-29">29 August 2010</date>
(air temperature = 26.2°C; 
<figureCitation id="134535E6FFF4FFB2FB5DFE814FCB0200" box="[1156,1221,295,319]" captionStart="FIGURE 6" captionStartId="11.[151,250,1784,1807]" captionTargetBox="[221,1371,238,1720]" captionTargetId="figure@11.[199,1388,227,1731]" captionTargetPageId="11" captionText="FIGURE 6. Spectrograms (above) and wave form (below) of advertisement call of (A) L. nigrops (KUHE 15430) from Kuala Lumpur, Malay Peninsula and (B) L. ingeri sp. nov. (KUHE 53834) from Santubong, Sarawak." httpUri="https://zenodo.org/record/281823/files/figure.png" pageId="12" pageNumber="34">Fig. 6</figureCitation>
B) had the duration from 305–763 (mean+SD = 485.9+243.8, n = 3/ 
<specimenCount id="9D78E2EAFFF4FFB2FD66FEEA4804025B" box="[703,778,332,356]" pageId="12" pageNumber="34" type="male">1 male</specimenCount>
) ms, and emitted intermittently. The call consisted of 5–9 (6.3+2.3, n = 3) short notes, each with the duration of 16–33 (20.7+5.1, n = 18) ms, and the note gap of 88–102 (93.7+3.8, n = 15) ms, with the note repetition rate of 11.8–16.4 (13.7+2.4, n = 3) per s. Each note is composed of 4–11 (6.1+2.0, n = 18) fine pulses. Frequency bands spread over 1.5 to 2.2 kHz range, with the dominant from 1,650 to 1,950 (1,792+75, n = 18) Hz. Each note showed weak frequency modulation, with the initial and middle frequencies (1,650 to 1,950 [1,792+75, n = 18] Hz) falling towards the end of the note (1,600–1,900 [1,760+99, n = 18] Hz).
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF1EFDE14BB10063" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
The calls recorded on 
<date id="FFC00FA3FFF4FFB2FE67FDE1499D0160" box="[446,659,583,607]" pageId="12" pageNumber="34" value="1990-12-26">26 December 1990</date>
at Niah (air temperature = 25.9°C) were basically similar in temporal structure (call duration 539–697 [618.2+111.7, n = 2/ 
<specimenCount id="9D78E2EAFFF4FFB2FD2BFDCA483201BB" box="[754,828,620,644]" pageId="12" pageNumber="34" type="male">1 male</specimenCount>
] ms; number of notes 8–10 [9.0, n = 2]; note duration 21–39 [25.8+5.1, n = 18] ms; note gap 72–105 [76.2+7.9, n = 16] ms; note repetition rate 14.3–18.4 [14.6, n = 2] per s; number of pulse per note 5–10 [7.0+1.5, n = 18]). However, frequencies were lower than in Santubong calls, with the dominant from 1,300 to 1,500 (1,392+62, n = 18) Hz. Lowering of the initial frequency (1,350 to 1,600 [1,450+132, n = 3] Hz) towards the end of the note (1,250–1,350 [1,300+41, n = 4] Hz) was similar to the Santubong calls. Three harmonic frequencies were present at approximately 1,875–1,930, 2,250–2,300, and 3,900–4,000 Hz.
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF1EFCC14BE10679" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
<emphasis id="B90AF571FFF4FFB2FF1EFCC14A6200BF" bold="true" box="[199,364,871,896]" pageId="12" pageNumber="34">Comparisons.</emphasis>
<taxonomicName id="4C7E52E0FFF4FFB2FEA0FCCE497A00BF" box="[377,628,872,896]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FEA0FCCE497A00BF" box="[377,628,872,896]" italics="true" pageId="12" pageNumber="34">Leptobrachium ingeri</emphasis>
</taxonomicName>
is placed in the 
<taxonomicName id="4C7E52E0FFF4FFB2FC91FCCF48CE00BF" box="[840,960,873,896]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FC91FCCF48CE00BF" box="[840,960,873,896]" italics="true" pageId="12" pageNumber="34">L. nigrops</emphasis>
</taxonomicName>
group by having small body size. This character distinguished it from all other species of 
<taxonomicName id="4C7E52E0FFF4FFB2FD15FC2A4879009B" box="[716,887,908,932]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="B90AF571FFF4FFB2FD15FC2A4879009B" box="[716,887,908,932]" italics="true" pageId="12" pageNumber="34">Leptobrachium</emphasis>
</taxonomicName>
, except for the other two species of the group (
<taxonomicName id="4C7E52E0FFF4FFB2FA5CFC2B4BE200F7" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FA5CFC2B4BE200F7" italics="true" pageId="12" pageNumber="34">L. nigrops</emphasis>
</taxonomicName>
and 
<taxonomicName id="4C7E52E0FFF4FFB2FEFDFC174AC300F7" box="[292,461,944,968]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kanowitense">
<emphasis id="B90AF571FFF4FFB2FEFDFC174AC300F7" box="[292,461,944,968]" italics="true" pageId="12" pageNumber="34">L. kanowitense</emphasis>
</taxonomicName>
). 
<taxonomicName id="4C7E52E0FFF4FFB2FE3CFC1649D500F7" box="[485,731,944,968]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FE3CFC1649D500F7" box="[485,731,944,968]" italics="true" pageId="12" pageNumber="34">Leptobrachium ingeri</emphasis>
</taxonomicName>
has relatively larger eye than 
<taxonomicName id="4C7E52E0FFF4FFB2FBEBFC174FAB00F7" box="[1074,1189,945,968]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FBEBFC174FAB00F7" box="[1074,1189,945,968]" italics="true" pageId="12" pageNumber="34">L. nigrops</emphasis>
</taxonomicName>
(15.0–18.5% SVL vs.12.0–17.3% SVL in 
<taxonomicName id="4C7E52E0FFF4FFB2FE4EFC73490000D3" box="[407,526,981,1004]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FE4EFC73490000D3" box="[407,526,981,1004]" italics="true" pageId="12" pageNumber="34">L. nigrops</emphasis>
</taxonomicName>
), and relatively larger values than 
<taxonomicName id="4C7E52E0FFF4FFB2FC6EFC734F6B00D3" box="[951,1125,980,1004]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kanowitense">
<emphasis id="B90AF571FFF4FFB2FC6EFC734F6B00D3" box="[951,1125,980,1004]" italics="true" pageId="12" pageNumber="34">L. kanowitense</emphasis>
</taxonomicName>
in snout to nostril length (7.9–10.4% SVL vs. 7.0–8.5% SVL in 
<taxonomicName id="4C7E52E0FFF4FFB2FD83FC5F480A072F" box="[602,772,1016,1040]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kanowitense">
<emphasis id="B90AF571FFF4FFB2FD83FC5F480A072F" box="[602,772,1016,1040]" italics="true" pageId="12" pageNumber="34">L. kanowitense</emphasis>
</taxonomicName>
), internarial distance (7.2–8.7% SVL vs. 6.2–7.0% SVL), intercanthal distance (17.6–20.7% SVL vs. 16.9–18.5% SVL), and inner metatarsal tubercle length (4.8–6.3% SVL vs. 4.3–4.4% SVL). In addition to morphometric differences, 
<taxonomicName id="4C7E52E0FFF4FFB2FCA0FBE748D10767" box="[889,991,1089,1112]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FCA0FBE748D10767" box="[889,991,1089,1112]" italics="true" pageId="12" pageNumber="34">L. ingeri</emphasis>
</taxonomicName>
differs from 
<taxonomicName id="4C7E52E0FFF4FFB2FB5CFBE74E3D0767" box="[1157,1331,1088,1112]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kanowitense">
<emphasis id="B90AF571FFF4FFB2FB5CFBE74E3D0767" box="[1157,1331,1088,1112]" italics="true" pageId="12" pageNumber="34">L. kanowitense</emphasis>
</taxonomicName>
in more developed toe webs: phalanges free of web 
<taxonomicName id="4C7E52E0FFF4FFB2FD5AFBCE49EA0740" box="[643,740,1128,1151]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FD5AFBCE499D0740" box="[643,659,1128,1151]" italics="true" pageId="12" pageNumber="34">L</emphasis>
. 
<emphasis id="B90AF571FFF4FFB2FD78FBCE49EA0740" box="[673,740,1128,1151]" italics="true" pageId="12" pageNumber="34">ingeri</emphasis>
</taxonomicName>
samples were 1–1 3/4 on the outer side of first (vs. 1 3/4–2 on 
<taxonomicName id="4C7E52E0FFF4FFB2FA5CFBCE4A2C0797" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kanowitense">
<emphasis id="B90AF571FFF4FFB2FA5CFBCE4A2C0797" italics="true" pageId="12" pageNumber="34">L. kanowitense</emphasis>
</taxonomicName>
), 1/4–2/3 on the outer side of second (vs. 1 1/4–1 3/4), 2 2/3-3 on the inner side of third (vs. 3–3 1/4), 1 1/4–1 3/4 on the outer side of third (vs. 2), 3 2/3 on the inner side of fourth (vs. 3 3/4–4), 3 2/3–3 3/4 on the outer side of fourth (vs. 3 3/4–4), and 1 1/2–1 2/3 on the inner side of fifth (vs. 2) toes. 
<taxonomicName id="4C7E52E0FFF4FFB2FCFEFB424F1307C3" box="[807,1053,1252,1276]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FCFEFB424F1307C3" box="[807,1053,1252,1276]" italics="true" pageId="12" pageNumber="34">Leptobrachium ingeri</emphasis>
</taxonomicName>
further differs from the other two species in much more sharp finger tips, and in more distinct markings on dorsum and ventrum than the other two species.
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF10FAF54E9C05B5" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
In calls, 
<taxonomicName id="4C7E52E0FFF4FFB2FEE9FAF54A9A0655" box="[304,404,1363,1386]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FEE9FAF54A9A0655" box="[304,404,1363,1386]" italics="true" pageId="12" pageNumber="34">L. ingeri</emphasis>
</taxonomicName>
closely resembles 
<taxonomicName id="4C7E52E0FFF4FFB2FDAAFAF549EC0655" box="[627,738,1363,1386]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FDAAFAF549EC0655" box="[627,738,1363,1386]" italics="true" pageId="12" pageNumber="34">L nigrops</emphasis>
</taxonomicName>
from Malay Peninsula: Calls of 
<taxonomicName id="4C7E52E0FFF4FFB2FBBCFAF54FD40655" box="[1125,1242,1363,1386]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="nigrops">
<emphasis id="B90AF571FFF4FFB2FBBCFAF54FD40655" box="[1125,1242,1363,1386]" italics="true" pageId="12" pageNumber="34">L. nigrops</emphasis>
</taxonomicName>
, recorded on 
<date id="FFC00FA3FFF4FFB2FA59FAF44A4506B1" pageId="12" pageNumber="34" value="1992-12-30">30 December 1992</date>
(air temperature = 26.4°C) and on 
<date id="FFC00FA3FFF4FFB2FD38FAD0489006B1" box="[737,926,1398,1423]" pageId="12" pageNumber="34" value="1993-01-21">21 January 1993</date>
(air temperature = 25.9°C) at the campus of University of 
<collectingCountry id="F36969F3FFF4FFB2FEEDFA3C4A85068D" box="[308,395,1434,1458]" name="Malaysia" pageId="12" pageNumber="34">Malaya</collectingCountry>
, Selangor, West 
<collectingCountry id="F36969F3FFF4FFB2FD9EFA3C49A3068D" box="[583,685,1434,1458]" name="Malaysia" pageId="12" pageNumber="34">Malaysia</collectingCountry>
were short trills (Fig. 11). The call varied in length from 618–851 (617.9+131.4, n = 3) ms and consisted of 8–12 (mean+SD = 9.7+2.1, n = 26/ 
<specimenCount id="9D78E2EAFFF4FFB2FC26FA184F5806E9" box="[1023,1110,1470,1494]" pageId="12" pageNumber="34" type="male">2 males</specimenCount>
) short notes, each with 4–26 (8.4+5.2, n = 29) pulses. The note duration varied from 14–85 (29.8+17.8, n = 29) ms, and the note gap from 72–111 (78.6+10.4, n = 26) ms. The note repetition rate varied from 12.9–14.3 (13.8+0.7, n = 3). Frequency band spread over 1,700 to 1,950 Hz range, with the dominant from 1,700 to 1,950 (1,852+75, n = 29) Hz. Within a note, the initial frequency (1,800–1,900 [1,840+39, n = 10] HZ) slightly lowered at the end (1,700–1,950 [1,777+82, n = 11] Hz). These characteristics overlap those of 
<taxonomicName id="4C7E52E0FFF4FFB2FD7FF9D5480905B5" box="[678,775,1651,1674]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FD7FF9D5480905B5" box="[678,775,1651,1674]" italics="true" pageId="12" pageNumber="34">L. ingeri</emphasis>
</taxonomicName>
, and the two species cannot be differentiated acoustically.
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF10F93048C105ED" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
<emphasis id="B90AF571FFF4FFB2FF10F9304A120591" bold="true" box="[201,284,1686,1710]" pageId="12" pageNumber="34">Range.</emphasis>
The coastal area of 
<collectingCountry id="F36969F3FFF4FFB2FDDBF930496C0591" box="[514,610,1686,1710]" name="Malaysia" pageId="12" pageNumber="34">Sarawak</collectingCountry>
in Malaysian Borneo (Santubong, Matang [Kuching Division], and Niah [Miri Division]); Belitung Island, 
<collectingCountry id="F36969F3FFF4FFB2FDCCF91C498C05ED" box="[533,642,1722,1746]" name="Indonesia" pageId="12" pageNumber="34">Indonesia</collectingCountry>
(Tanjung Padang) (
<figureCitation id="134535E6FFF4FFB2FC86F91C48B305ED" box="[863,957,1722,1746]" captionStart="FIGURE 1" captionStartId="1.[151,250,1191,1214]" captionTargetBox="[170,1416,368,1148]" captionTargetId="figure@1.[151,1436,359,1156]" captionTargetPageId="1" captionText="FIGURE 1. Map of Malaysia and western part of Indonesia, showing sample localities of the L. nigrops group that are used in the molecular analyses (triangle) and morphological analyses (reversed triangle). Blue: L. nigrops. Yellow: L. ingeri sp. nov. Red: L. kanowitense sp. nov.). Details of localities are shown in Table 1." httpUri="https://zenodo.org/record/281818/files/figure.png" pageId="12" pageNumber="34">Figure 1</figureCitation>
).
</paragraph>
<paragraph id="8BC12963FFF4FFB2FF10F978482804CD" blockId="12.[151,1437,151,2034]" pageId="12" pageNumber="34">
<emphasis id="B90AF571FFF4FFB2FF10F9784A8505C8" bold="true" box="[201,395,1758,1783]" pageId="12" pageNumber="34">Natural history.</emphasis>
Larval, and other ecological data are lacking, but breeding season seems to include late August and late December, as evidenced by calling males. Matang, Santubong, and Tanjung Padang, where the 
<typeStatus id="54C597C1FFF4FFB2FF4EF8814BC90400" box="[151,199,1831,1855]" pageId="12" pageNumber="34">type</typeStatus>
series of 
<taxonomicName id="4C7E52E0FFF4FFB2FEECF8814A980401" box="[309,406,1831,1854]" class="Amphibia" family="Megophryidae" genus="Leptobrachium" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="ingeri">
<emphasis id="B90AF571FFF4FFB2FEECF8814A980401" box="[309,406,1831,1854]" italics="true" pageId="12" pageNumber="34">L. ingeri</emphasis>
</taxonomicName>
were collected, are lowland (&lt;
<quantity id="4C868486FFF4FFB2FD2BF88048230401" box="[754,813,1830,1854]" metricMagnitude="1" metricUnit="m" metricValue="6.0" pageId="12" pageNumber="34" unit="m" value="60.0">60 m</quantity>
a.s.l.). The 
<typeStatus id="54C597C1FFF4FFB2FC6DF8804F180401" box="[948,1046,1830,1854]" pageId="12" pageNumber="34" type="holotype">holotype</typeStatus>
was collected under a dead tree in an old trail near a small river in secondary forest. Other species found on this trail were: 
<taxonomicName id="4C7E52E0FFF4FFB2FB1AF8ED4A0B04B9" authority="Tschudi" authorityName="Tschudi" class="Amphibia" family="Dicroglossidae" genus="Limnonectes" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kuhlii">
<emphasis id="B90AF571FFF4FFB2FB1AF8ED4E92045D" box="[1219,1436,1866,1890]" italics="true" pageId="12" pageNumber="34">Limnonectes kuhlii</emphasis>
(Tschudi)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FEC9F8C9499C04B9" authority="Andersson" authorityName="Andersson" box="[272,658,1902,1926]" class="Amphibia" family="Dicroglossidae" genus="Limnonectes" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="leporinus">
<emphasis id="B90AF571FFF4FFB2FEC9F8C9490104B9" box="[272,527,1902,1926]" italics="true" pageId="12" pageNumber="34">Limnonectes leporinus</emphasis>
Andersson
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FD44F8C94F1704B9" authority="Inger" authorityName="Inger" box="[669,1049,1902,1926]" class="Amphibia" family="Dicroglossidae" genus="Limnonectes" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="kenepaiensis">
<emphasis id="B90AF571FFF4FFB2FD44F8C948CF04B9" box="[669,961,1902,1926]" italics="true" pageId="12" pageNumber="34">Limnonectes kenepaiensis</emphasis>
(Inger)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FBFCF8C94BD60495" authority="Gravenhorst" authorityName="Gravenhorst" class="Amphibia" family="Dicroglossidae" genus="Fejervarya" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="limnocharis">
<emphasis id="B90AF571FFF4FFB2FBFCF8C94E3E04B9" box="[1061,1328,1902,1926]" italics="true" pageId="12" pageNumber="34">Fejervarya limnocharis</emphasis>
(Gravenhorst)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FF31F83449280495" authority="Gunther" authorityName="Gunther" box="[232,550,1938,1962]" class="Amphibia" family="Bufonidae" genus="Ansonia" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="leptopus">
<emphasis id="B90AF571FFF4FFB2FF31F8344AA20495" box="[232,428,1938,1962]" italics="true" pageId="12" pageNumber="34">Ansonia leptopus</emphasis>
(Günther)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FDEBF835489B0495" authority="Gunther" authorityName="Gunther" box="[562,917,1938,1962]" class="Amphibia" family="Ranidae" genus="Meristogenys" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="jerboa">
<emphasis id="B90AF571FFF4FFB2FDEBF83548150495" box="[562,795,1938,1962]" italics="true" pageId="12" pageNumber="34">Meristogenys jerboa</emphasis>
(Günther)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FC7BF8344FED0495" authority="Gunther" authorityName="Gunther" box="[930,1251,1938,1962]" class="Amphibia" family="Ranidae" genus="Hylarana" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="signata">
<emphasis id="B90AF571FFF4FFB2FC7BF8344F660495" box="[930,1128,1938,1962]" italics="true" pageId="12" pageNumber="34">Hylarana signata</emphasis>
(Günther)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FB29F8344A3404F1" authority="Peters" authorityName="Peters" class="Amphibia" family="Ranidae" genus="Hylarana" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="raniceps">
<emphasis id="B90AF571FFF4FFB2FB29F8344BC704F1" italics="true" pageId="12" pageNumber="34">Hylarana raniceps</emphasis>
(Peters)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FE8CF810483904F1" authority="Gravenhorst" authorityName="Gravenhorst" box="[341,823,1974,1998]" class="Amphibia" family="Rhacophoridae" genus="Polypedates" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="leucomystax">
<emphasis id="B90AF571FFF4FFB2FE8CF810498E04F1" box="[341,640,1974,1998]" italics="true" pageId="12" pageNumber="34">Polypedates leucomystax</emphasis>
(Gravenhorst)
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FC8BF8104FE104F1" authority="Boulenger" authorityName="Boulenger" box="[850,1263,1974,1998]" class="Diplopoda" family="Chelodesmidae" genus="Rhacophorus" kingdom="Animalia" order="Polydesmida" pageId="12" pageNumber="34" phylum="Arthropoda" rank="species" species="fasciatus">
<emphasis id="B90AF571FFF4FFB2FC8BF8104F6D04F1" box="[850,1123,1974,1998]" italics="true" pageId="12" pageNumber="34">Rhacophorus fasciatus</emphasis>
Boulenger
</taxonomicName>
, 
<taxonomicName id="4C7E52E0FFF4FFB2FAD2F8104AAF04CD" authority="Boulenger" authorityName="Boulenger" class="Amphibia" family="Microhylidae" genus="Kalophrynus" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="heterochirus">
<emphasis id="B90AF571FFF4FFB2FAD2F8104A2D04CD" italics="true" pageId="12" pageNumber="34">Kalophrynus heterochirus</emphasis>
Boulenger
</taxonomicName>
, and 
<taxonomicName id="4C7E52E0FFF4FFB2FE02F87C482804CD" authority="Parker." authorityName="Parker." box="[475,806,2010,2034]" class="Amphibia" family="Microhylidae" genus="Microhyla" kingdom="Animalia" order="Anura" pageId="12" pageNumber="34" phylum="Chordata" rank="species" species="borneensis">
<emphasis id="B90AF571FFF4FFB2FE02F87C49C104CD" box="[475,719,2010,2034]" italics="true" pageId="12" pageNumber="34">Microhyla borneensis</emphasis>
Parker.
</taxonomicName>
</paragraph>
</subSubSection>
</treatment>
</document>